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. 2016 Jul 22;444(2):1515–1541. doi: 10.1016/j.jmaa.2016.07.028

Asymptotic profile in selection–mutation equations: Gauss versus Cauchy distributions

Àngel Calsina a, Sílvia Cuadrado a,, Laurent Desvillettes b, Gaël Raoul c
PMCID: PMC7094311  PMID: 32226135

Abstract

In this paper, we study the asymptotic (large time) behaviour of a selection–mutation–competition model for a population structured with respect to a phenotypic trait when the rate of mutation is very small. We assume that the reproduction is asexual, and that the mutations can be described by a linear integral operator. We are interested in the interplay between the time variable t and the rate ε of mutations. We show that depending on α>0, the limit ε0 with t=εα can lead to population number densities which are either Gaussian-like (when α is small) or Cauchy-like (when α is large).

Keywords: Selection–mutation equations, Asymptotic behaviour, Spectral theory, Population dynamics

1. Introduction

1.1. Selection–mutation–competition models

The phenotypic diversity of a species impacts its ability to evolve. In particular, the importance of the variance of the population along a phenotypic trait is illustrated by the fundamental theorem of natural selection [15], and the breeder's equation [21]: the evolution speed of a population along a one dimensional fitness gradient (or under artificial selection) is proportional to the variance of the initial population. Recently, the phenotypic variance of populations has also come to light as an important element to describe the evolutionary dynamics of ecosystems (where many interacting species are considered) [27], [4], [26].

Over the last decade, the issue of Evolutionary Rescue emerged as an important question [3], [10], [17] (see also the seminal work of Luria and Delbrück [20]), and led to a new interest in the phenotypic distribution of populations, beyond phenotypic variance. Evolutionary Rescue is concerned with a population living in an environment that changes suddenly. The population will survive either if some individuals in the population carry an unusual trait that turns out to be successful in the new environment, or if new mutants able to survive in the new environment appear before the population goes extinct (see [22] for a discussion on the relative effect of de novo mutations and standing variance in Evolutionary Rescue). In any case, the fate of the population will not be decided by the properties of the bulk of its density, but rather by the properties of the tail of the initial distribution of the individuals, close to the favourable traits for the new environment. A first example of such problems comes from emerging diseases [16]: Animal infections sometimes are able to infect humans. This phenomenon, called zoonose, is the source of many human epidemics: HIV, SARS, Ebola, MERS-CoV, etc. A zoonose may happen if a pathogen that reaches a human has the unusual property of being adapted to this new human host. A second example comes from the emergence of microbes resistant to an antimicrobial drug that is suddenly spread in the environment of the microbe. This second phenomenon can easily be tested experimentally [3], [24], and has major public health implications [9].

Most papers devoted to the genetic diversity of populations structured by a continuous phenotypic trait describe the properties of mutation–selection equilibria. It is however also interesting to describe the genetic diversity of populations that are not at equilibrium (transient dynamics): pathogen populations for instance are often in transient situations, either invading a new host, or being eliminated by the immune system. We refer to [18] for a review on transient dynamics in ecology. For asexual populations structured by a continuous phenotypic trait, several models exist, corresponding to different biological assumptions [11]. If the mutations are modelled by a diffusion, the steady populations (for a model close to (1), but where mutations are modelled by a Laplacian) are Gaussian distributions [19], [6]. Furthermore, [1], [14] have considered some transient dynamics for this model. In the model that we will consider (see (1)), the mutations are modelled by a non-local term. It was shown in [7] (see also [6]) that mutation–selection equilibria are then Cauchy profiles (under some assumptions), and this result has been extended to more general mutation kernels in [8], provided that the mutation rate is small enough. Finally, let us notice that the case of sexual populations is rather different, since recombinations by themselves can imply that a mutation–recombination equilibrium exists, even without selection. We refer to the infinitesimal model [5], and to [25] for some studies on the phenotypic distribution of sexual species in a context close to the one presented here for asexual populations.

In this article, we consider a population consisting of individuals structured by a quantitative phenotypic trait xI (I open interval of R containing 0), and denote by f:=f(t,x)0 its density. Here, the trait x is fully inherited by the offspring (if no mutation occurs), so that x is indeed rather a breeding value than a phenotypic trait (see [23]). We assume that individuals reproduce with a rate 1, and die at a rate

x2+If(t,y)dy.

This means that individuals with trait x=0 are those who are best adapted to their environment, and that the fitness decreases like a parabola around this optimal trait (this is expected in the surroundings of a trait of maximal fitness). It also means that the strength of the competition modelled by the logistic term is identical for all traits. When an individual of trait xI gives birth, we assume that the offspring will have the trait x with probability 1ε, and a different trait x with probability ε(0,1). ε is then the probability that a mutation affects the phenotypic trait of the offspring. We can now define the growth rate of the population of trait x (that is the difference between the rate of births without mutation, minus the death rate) as

rε(t,x)=1εx2If(t,y)dy.

When a mutation affects the trait of the offspring, we assume that the trait x of the mutated offspring is drawn from a law over the set of phenotypes IR with a density γ:=γ(x)L1(I). The function γ then satisfies

γ(x)0,Iγ(x)dx=1

(we assume moreover in some of the mathematical statements that γ is bounded, C1, with bounded derivative and that it is strictly positive on I). The main assumption here is that the law of the trait of a mutated offspring does not depend on the trait of its parent. This classical assumption, known as house of cards is not the most realistic, but it can be justified when the mutation rate is small [6] (see also [8]). All in all, we end up with the following equation:

fε(t,x)t=rε(t,x)fε(t,x)+εγ(x)Ifε(t,y)dy. (1)

This paper is devoted to the study of the asymptotic behaviour of the solutions of equation (1) when ε is small and t large and it is organized as follows. In the rest of Section 1 the main results are quoted, first in an informal way, and then as rigorous statements. Section 2 contains the proof of Proposition 1.1 and Theorem 1.2, and finally, in Section 3, Theorem 1.3 is proved.

1.2. Asymptotic study of the model

When we consider the solutions of (1), two particular profiles naturally appear:

  • A Cauchy profile: For a given mutation rate ε>0 small enough, one expects that fε(t,x) will converge, as t goes to infinity, to the unique steady-state of (1), which is the following Cauchy profile
    fε(,x):=εγ(x)Iε()Iε()(1ε)+x2, (2)
    where Iε() is such that Ifε(,x)dx=Iε(). This steady-state of (1) is the so-called mutation–selection equilibrium of the House of cards model (1), which has been introduced in [7] (we also refer to [6] for a broader presentation of existing results).
  • A Gaussian profile: If ε=0, the solution of (1) can be written as
    f0(t,x)=f(0,x)e0tI0(s)ds+ttx2, (3)
    where I0(t):=If0(t,x)dx, so that a Gaussian-like behaviour (with respect to x) naturally appears in this case. Surprisingly, we are not aware of any reference to this property in the population genetics literature.

We will show that, as suggested by the above arguments, we can describe the phenotypic distribution of the population, that is xfε(t,x), when either t1 (large time for a given mutation rate ε>0), or 0ε1 (small mutation rate, for a given time interval t[0,T]). Before providing the precise statements of our results (see Subsection 1.3), we will briefly describe them here, and illustrate them with numerical simulations. The numerical simulations presented in Fig. 1 and Fig. 2 are obtained thanks to a finite difference scheme (explicit Runge–Kutta in time), and we illustrate our result with a single simulation of (1) with ε=102, I=[3/2,3/2], γ(x)=140πex220 and fε(0,x)=Γ2(ε,x1) (see the definition of Γ2 in eq. (4) below). The initial condition corresponds to a population at the mutation–selection equilibrium which environment suddenly changes (the optimal trait originally in x=1 moves to x=0 at t=0). This example is guided by the Evolutionary Rescue experiments described in Subsection 1.1, where the sudden change is obtained by the addition of e.g. salt or antibiotic to a bacterial culture.

Fig. 1.

Fig. 1

The different graphs correspond to different time points, from t = 0 to t = 175 000, of the same simulation of (1) for ε = 10−2 (see in the text for a complete description). In each of these plots, the blue (resp. red, black) line represents xfε(t,x) (resp. x↦Γ1(t,ε,x), x↦Γ2(ε,x)). Note that in this figure, the scales of both axis change from one graph to the other, to accommodate with the dynamics of the solution f(t,⋅). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

Fig. 2.

Fig. 2

Simulation of (1) with ε = 10−2 (see in the text for a complete description). The red line represents fε(t,)Γ1(t,ε,)L1(I), while the black line represents fε(t,)Γ2(ε,)L1(I). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

We describe two phases of the dynamics of the population:

  • Large time: Cauchy profile. We show that fε(t,x) is asymptotically (when the mutation rate ε>0 is small) close to
    Γ2(ε,x)=εγ(0)γ(0)2π2ε2+x2, (4)
    provided tε4. The population is then a time-independent Cauchy distribution for large times. This theoretical result is coherent with numerical results: we see in Fig. 1 that fε(t,) is well described by Γ2(ε,), as soon as t105, which is confirmed by the value of fε(t,)Γ2(ε,)L1(I) for t105 given by Fig. 2.
  • Short time: Gaussian profile. We also show that fε(t,x) is asymptotically (when the mutation rate ε>0 is small) close to
    Γ1(t,ε,x)=f(0,x)tf(0,0)Iex2dxex2t, (5)
    provided 1tε2/3. The population has then a Gaussian-type distribution for short (but not too short) times. This theoretical result is coherent with numerical simulations: we see in Fig. 1 that fε(t,) is well described by Γ1(t,ε,) for t[102,104], which is confirmed by the value of fε(t,)Γ2(ε,)L1(I) for t[102,104] given by Fig. 2.

Another way to look at these results is to consider t0 and ε>0 as two parameters, and to see the approximations presented above as approximations of fε(t,) for some set of parameters: fε(t,)ε0Γ2(ε,) for (t,ε){(t˜,ε˜);t˜ε˜4}, while fε(t,)ε0Γ1(t,ε,) for (t,ε){(t˜,ε˜);1t˜ε˜2/3}. We have represented these sets in Fig. 3 .

Fig. 3.

Fig. 3

Representation of the set {(t˜,ε˜);t˜ε˜4} (in blue), where the approximation fε(t,⋅)∼ε → 0Γ2(ε,⋅) holds provided that ε > 0 is small enough; and of the set {(t˜,ε˜);1t˜ε˜2/3} (in red), where the approximation fε(t,⋅)∼ε → 0Γ1(t,ε,⋅) holds provided that ε > 0 is small enough. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

As described in the Subsection 1.1, the phenotypic distribution of species is involved in many ecological and epidemiological problematics. Our study is a general analysis of this problem and we do not have a particular application in mind. An interesting and (to our knowledge) new feature described by our study is that the tails of the trait distribution in a population can change drastically between “short times”, that is 1tε2/3 and “large times”, that is tε4: the distribution is initially close to a Gaussian distribution, with small tails, and then converges to a thick tailed Cauchy distribution. This result could have significant consequences for evolutionary rescue: the tails of the distribution then play an important role. Quantifying the effect of this property of the tails of the distributions would however require further work, in particular on the impact of stochasticity (the number of pathogen is typically large, but finite). The plasticity of the pathogen (see [12]) may also play an important role.

1.3. Main (rigorous) statements

Here we state the two main theorems of the paper, together with a proposition asserting the well-posedness of the equation that we consider.

We start with the issue of well-posedness:

Proposition 1.1

We consider the interval I ( I=]a,b[ , a<b+ ). We assume that γ is continuous, bounded and strictly positive on I, and that Iγ(x)dx=1 . We consider an initial datum f0 which is nonnegative, non-identically 0, and integrable on I. Finally we take ε]0,1/2[ .

Then the initial value problem (1) (with f(0,)=f0 ) admits a unique nonnegative (global in time) mild solution in C(R+;L1(I)) .

Then we turn to the two statements which are relative to the asymptotic behaviour when ε0 and t+.

Theorem 1.2

Under the assumptions of Proposition 1.1 , and assuming moreover that γ has a bounded derivative on I, there exist positive constants K,Kˆ and K˜ (independent of ε) such that the mild solution f of (1) satisfies (for t>0 )

xf(t,x)εγ(0)(γ(0)πε)2+x2L1(I)KεKˆε2eKˆε2t+K˜εln(1ε).

Theorem 1.3

Under the assumptions of Proposition 1.1 , and assuming moreover that a and b are finite, f0W1,(I) , f0(0)>0 and If0(x)dx<1 , there exists a constant C>0 (independent of ε) such that the mild solution f of (1) satisfies (for t>4 )

xf(t,x)f(0,x)tex2tf(0,0)Iey2dyL1(I)C(1t+εt32)eCεt. (6)

A careful check of the proof shows that the constant C appearing in (6) only depends on |ab| , some upper bounds on γL and f0W1, , and a lower bound on f0(0) .

Note first that, as previously stated in the introduction, if we consider times t which are large enough, for example t=ε4δ with δ>0, then, since (ε2lnεε2δ) is asymptotically equivalent to ε2δ, Theorem 1.2 ensures that

xf(ε4δ,x)εγ(0)(γ(0)πε)2+x2L1(I)=Oε0(εln(1ε)),

so that the Cauchy profile is indeed asymptotically reached for very large times.

The interpretation of Theorem 1.3 is a little more intricate, since it concerns intermediate times. For example, it implies the following description of the population's phenotypic diversity during transitory times, that is times t satisfying 1tε23:

There exists C>0 such that for κ>0 and ε>0 small enough (in such a way that the interval [κ2,κ23ε23] is not empty, that is ε<κ4),

t[κ2,κ23ε23],xf(t,x)f(0,x)tex2tf(0,0)Iey2dyL1(I)Cκ.

For example, if t=ε1/2, then

xf(t,x)f(0,x)tex2tf(0,0)Iey2dyL1(I)Cε1/4=Ct1/2.

Note finally that these results hold for models which are slightly more general than equation (1). In fact, in both theorems one can assume that the competition term is a weighted population instead of the total population number. In Theorem 1.3, one could also assume that the mutation kernel depends on the parents trait.

2. Proof of Proposition 1.1 and Theorem 1.2

We start here the proofs of Proposition 1.1 and of Theorem 1.2. We recall that

I=]a,b[, a<0<b, and γ:=γ(x) is a bounded, continuous, strictly positive function, such that γ(x)>0 and Iγ(x)dx=1. For the proof of Theorem 1.2, we will also assume that it has a bounded derivative on I.

We begin with the study of the linear operator associated to eq. (1).

2.1. Spectrum of the linear operator

Let us define

(Aεf)(x):=(1ε)f(x)x2f(x)+εγ(x)If(y)dy (7)

as the operator corresponding to the linear part in eq. (1).

We begin with a basic lemma which enables to define the semigroup generated by this operator.

Lemma 2.1

The linear operator Aε , defined on L1(I) and with domain D(Aε)={fL1(I):Ix2|f(x)|dx<} , generates an irreducible positive C0 -semigroup (denoted from now on by Tε(t) ).

Proof

The multiplication linear operator (Aε0f)(x):=(1ε)f(x)x2f(x) is the generator of a positive C0-semigroup.

Since γ is strictly positive, AεAε0 is a positive bounded perturbation whose only invariant closed ideals are 0 and the whole space L1(I). So Tε(t) is irreducible (see [13], Corollary 9.22). □

Next, we present a proposition which gives information about the spectrum of Aε.

Proposition 2.2

The linear operator Aε has only one eigenvalue. It is a strictly dominant algebraically simple eigenvalue λε>1ε and a pole of the resolvent, with corresponding normalized positive eigenvector

ψε(x)=εγ(x)λε(1εx2).

Moreover, for ε small enough, λε<1 .

The rest of the spectrum of the linear operator Aε is equal to the interval J=[min(1εa2,1εb2),1ε] .

Proof

In the sequel, the norm is the L1 norm on I. Let us first show that any λ belonging to the set J=Range(1εx2) belongs to the spectrum of Aε. In order to do this, for λ=1εx02,x0I˚, let us define fn(x)=n2(χ[x0,x0+1n](x)χ[x01n,x0](x)) for n such that [x01n,x0+1n]I. We then have fn=1 and (AελId)fn=n2x01nx0+1n|x2x02|dx0. So [min(1εa2,1εb2),1ε] is contained in the spectrum of Aε. The claim follows from the fact that the spectrum is a closed set.

On the other hand, notice that (for x0I), 1εx02 is not an eigenvalue, since the potential corresponding eigenfunction γ(x)x02x2 is not an integrable function on I (remember that γ does not vanish).

Let us now compute the resolvent operator of Aε, that is, let us try to solve the equation

Aεfλf=gL1(I). (8)

For λJ, defining p:=If(y)dy, (8) gives

f(x)=εγ(x)pg(x)λ(1εx2). (9)

Integrating, we get

(1εIγ(x)λ(1εx2)dx)p=Ig(x)λ(1εx2)dx, (10)

and λ belongs to the resolvent set unless the factor of p on the left hand side vanishes. Therefore σ(A)=J{λC:εIγ(x)λ(1εx2)dx=1}.

Since for any real number λ>1ε, the function Fε(λ):=εIγ(x)λ(1εx2)dx is continuous, strictly decreasing, and satisfies limλ1εFε(λ)=+ (recall that γ(0)>0) and limλ+Fε(λ)=0, we see that there is a unique real solution of Fε(λ)=1 in (1ε,). We denote it by λε.

Taking g(x)=0 in (8), we see that λε is an eigenvalue with corresponding normalized strictly positive eigenvector

ψε=εγ(x)λε(1εx2).

Taking g=ψε and λ=λε we see that the left hand side in (10) vanishes, whereas the right hand side is strictly negative, so that Aεfλεf=ψε has no solution and hence λε is algebraically simple.

Indeed, it also follows from (10) that the range of AελεId coincides with the kernel of the linear form defined on L1(I) by the L function 1λε(1ε)+x2 (which is the eigenvector corresponding to the eigenvalue λε of the adjoint operator Aε) and hence it is a closed subspace of L1(I). Therefore, λε is a pole of the resolvent (see Theorem A.3.3 of [13]). Furthermore, since

Fε(1)=εIγ(x)ε+x2dx=Iγ(x)1+(xε)2dxε00,

we see that Fε(1)<1 for ε small enough, and hence λε<1.

Substituting λ by a+bi in the characteristic equation

1+εIγ(x)(1εx2λ)dx=0, (11)

we have that the imaginary part is εbIγ(x)(1εx2λ)dx. Since γ(x)>0, there are no nonreal solutions of (11). □

Remark 2.1

Note that limε0λε=1.

We now write an expansion of the eigenvalue λε.

Proposition 2.3

Let us assume that γ has a bounded derivative on I. Let λε be the dominant eigenvalue of the operator Aε . Then

|λε(1ε)γ(0)2π2ε2|=Oε0(ε3ln1ε)

Proof

Let us consider the change of variable x=νεz where νε=λε(1ε). We have

1=εabγ(x)(λε(1εx2))dx=εaνεbνεγ(νεz)νε2+(νεz)2νεdz=ενεaνεbνεγ(νεz)1+z2dz.

Then

|νεεγ(0)π|=|aνεbνεγ(νεz)1+z2dzγ(0)π||aνεbνεγ(νεz)1+z2dzRγ(νεz)1+z2dz|+|Rγ(νεz)γ(0))1+z2dz|4γBνε+dz1+z2+2γνε0Aνεz1+z2dz,

where we have used

|γ(νεz)γ(0)|min(γ,γνε|z|),

and have denoted A:=γγ and B:=min(|a|,b,A).

Since

4γBνε+dz1+z2=4γarctan(νεB)4γνεB,

and

2γνε0Aνεz1+z2dz=γνεln(1+A2νε2),

we obtain

|νεεγ(0)π|ενε(4γB+γln(1+A2νε2)), (12)

which implies

ε(γ(0)πνε(4γB+γln(1+A2νε2)))νεε(γ(0)π+νε(4γB+γln(1+A2νε2))).

Since

νε(4γB+γln(1+A2νε2))ε00,

we have

γ(0)πε2νε2γ(0)πε (13)

for ε small enough.

Therefore, using (13) in (12) we get

|νεεγ(0)π|ε22γ(0)π(4γB+γln(1+4A2γ(0)2π2ε2))Cε2ln(1ε). (14)

Finally, by (13) and (14),

|λε(1ε)γ(0)2π2ε2|=|νε+γ(0)πε||νεγ(0)πε|3γ(0)πCε3ln(1ε).

 □

2.2. Proof of Proposition 1.1

Let us define A˜ε=AελεId and let T˜ε(t)=eλεtTε(t) be the semigroup generated by A˜ε. We now rewrite equation (1) as

f(t,x)t=A˜εf(t,x)+(λεIf(t,y)dy)f(t,x). (15)

We look for solutions of (15) (with positive initial condition f0L1(I)) which can be written as f(t,x)=h(t)(T˜ε(t)f0)(x), with h:=h(t) a real valued function of time such that h(0)=1. Substituting in (15), it follows that f is indeed a solution of eq. (1) if h(t) satisfies the following initial value problem for an ordinary differential equation:

h(t)=(λεh(t)I(T˜ε(t)f0)(x)dx)h(t),h(0)=1. (16)

Then h satisfies the integral equation

h(t)=1+0t(λεh(s)I(T˜εf0)(x)dx)h(s)ds,

from which the following identity follows

h(t)T˜ε(t)f0=T˜ε(t)f0+0tT˜ε(ts)(λεh(s)I(T˜ε(s)f0)(x)dx)h(s)T˜ε(s)f0ds,

i.e., f(t,x) is a solution of the variations of constants equation.

On the other hand, the nonlinear part of the right hand side of (15) is a locally Lipschitz function of fL1(I). From this uniqueness follows, whereas global existence and nonnegativity are clear from (16).

2.3. Asymptotic behaviour of the nonlinear equation

Let us start this subsection with a proposition ensuring in a quantitative way the convergence of the solution towards a nontrivial steady state:

Proposition 2.4

Under the assumptions of Theorem 1.2 , for ε>0 small enough, and any ρε<(γ(0)πε)2 , there exists a constant Cε>0 (depending on f0 and ε) such that

f(t,)λεψεL1(I)Cεeρεt.

Furthermore, taking ρε=αε2=λε(1ε)2 , the following more explicit (in terms of dependence w.r.t ε) estimate holds

f(.,t)λεψεL1(I)KεKˆε2eαεt2,

where K,Kˆ>0 depend on f0 but not on ε.

The rest of this subsection will be devoted to proving Proposition 2.4 taking advantage of the special form of the solution, f(t)=h(t)Tε(t)f0, as written in the proof of Proposition 1.1. In order to do so, we will also use some lemmas (Lemma 2.5, Lemma 2.6, Lemma 2.7, Lemma 2.8, Lemma 2.9, Lemma 2.10 which are stated below).

Let us start with one in which some properties of the spectrum of A˜ε=AελεId are used to study the asymptotic behaviour of the semigroup T˜ε(t) generated by A˜ε.

Lemma 2.5

  • a)

    The essential growth bound of the semigroup generated by A˜ε is ωess(T˜ε)=1ελε .

  • b)

    The growth bound of the semigroup generated by A˜ε is ω0(T˜ε)=0 .

Proof

  • a)

    A˜ε is a compact (one rank) perturbation of A˜ε0f:=(1εx2λε)f. Then ωess(T˜ε)=ωess(T˜ε0) where T˜ε0(t) is the semigroup generated by A˜ε0 (see [2]).

    Since A˜ε0 is a multiplication operator, ωess(T˜ε0)=1ελε and the result follows.

  • b)

    By Proposition 2.2, the spectral bound of A˜ε is 0 and the spectral mapping theorem holds for any positive C0-semigroup on L1 (see [13]).  □

Let us now write, for a positive non-identically zero f0, (T˜ε(t))f0(x)=cf0ψε(x)+v(t,x) where ψε(x) is the eigenvector corresponding to the eigenvalue 0 of A˜ε, and cf0ψε(x) is the spectral projection of f0 on the kernel of A˜ε (Note that cf0>0 since f0 is positive and A˜ε is the generator of an irreducible positive semigroup.) We also define φ(t):=Iv(t,x)dx.

The following lemma gives the asymptotic behaviour of cf0:

Lemma 2.6

Let us assume that f0 is a positive integrable function on I. Then there exist positive constants K1 , K2 (independent of ε but depending on f0 ) such that K1ε2cf0K2 . Moreover, limε0cf0=0 .

Proof

Recall that cf0=ψε,f0 where ψε is the eigenvector of the adjoint operator Aε corresponding to the eigenvalue λε, normalized such that ψε,ψε=1. Since

ψε=(εIγ(x)(λε(1εx2))2dx)1λε(1εx2),

we see that

cf0=If0(x)λε(1εx2)dxεIγ(x)(λε(1εx2))2dx.

Let us start by bounding the denominator from above. Using that, by Proposition 2.3, for ε small enough, λε(1ε)(γ(0)πε)22, we obtain the bound

εIγ(x)(λε(1εx2))2dxεsupxγ(x)R1((γ(0)πε)22+x2)2dx=supxγ(x)2γ(0)3(πε)2=:K0ε2. (17)

Similarly, since for ε small enough, λε(1ε)2(γ(0)πε)2, we have

εIγ(x)(λε(1εx2))2dxεmin[x0,x0]γ(x)x0x0dx(2(γ(0)πε)2+x2)2K3ε2, (18)

where x0 is any positive value in I.

For the numerator we have, on the one hand,

ε2If0(x)λε(1εx2)dxIε2(γ(0)πε)22+x2f0(x)dx, (19)

where the right hand side tends to 0 when ε goes to 0 by an easy application of the Lebesgue dominated convergence theorem (note that the integrand is bounded above by 2(γ(0)π)2f0(x)).

On the other hand, notice that there exists an interval JI which does not contain 0 such that Jf0(x)dx>0. Then, since

If0(x)λε(1εx2)dxJf0(x)λε(1εx2)dx

and

limε0Jf0(x)λε(1εx2)dx=Jf0(x)x2dx>0,

there exists a constant K4>0 such that

If0(x)λε(1εx2)dx>K4. (20)

By (18) and (19),

cf0=ε2If0(x)λε(1εx2)dxε3Iγ(x)λε(1εx2)dxε2If0(x)λε(1εx2)dxK3ε00

and by (17) and (20), and ε small enough,

cf0K4K0ε2=:K1ε2.

This completes the proof. □

Remark 2.2

If f0(x) is bounded below by a positive number c in a neighbourhood (δ,δ) of 0, then the lower estimate can be improved using that

δδεk2ε2+x2dx=2karctan(δkε)ε0+πk.

Indeed, for ε small enough

εIf0(x)λε(1εx2)dxεIf0(x)2(γ(0)πε)2+x2dxcδδε(2γ(0)π)2ε2+x2dxε0+c2γ(0).

So in this case, for ε small enough,

cf0c2γ(0)εK0ε2=:Kε

for some constant K independent of ε.

The next two lemmas enable to estimate φ(t) (defined immediately before the statement of Lemma 2.6). In the first one, the dependence w.r.t. ε is not explicit.

Lemma 2.7

For ε small enough and any ρε<(γ(0)πε)2 there exists Kε>0 such that |φ(t)|v(t,)Kεeρεtf0 .

Proof

Since, by Lemma 2.5, ωess(A˜ε)<ω0(A˜ε), we can apply Theorem 9.11 in [13], and get the estimate

v(t,)=T˜ε(t)f0cf0ψεKεeηtf0η<λε(1ε).

Proposition 2.3 gives then the statement. □

We now give an estimate of the dependence of Kε on ε, provided that ρε is chosen far enough from its limit value. More precisely, we choose ρε=λε(1ε)2=:αε2.

Lemma 2.8

For ε small enough, there exists a constant K independent of ε and of f0 such that

T˜ε(t)f0cf0ψεKε4eαε2tf0.

Proof

Since the proof of this result is quite technical, we delay it to the end of this section (subsection 2.5). □

We now turn to the study of the scalar function h(t). Notice that (16) can be written as

h(t)=(λε(cf0+φ(t))h(t))h(t),h(0)=1. (21)

The next two lemmas are devoted to the analysis of the asymptotic behaviour of h(t). In the first one, the dependence w.r.t. ε of the constants is not explicit.

Lemma 2.9

For ε>0 small enough and any ρε<(γ(0)πε)2 , there exists a positive constant Cˆε>0 such that |h(t)λεcf0|Cˆεeρεt .

Proof

The solution of (21) is explicitly given by

h(t)=eλεt1+0t(cf0+φ(s))eλεsds=1eλεt+cf0λε(1eλεt)+eλεt0tφ(s)eλεsds.

Then

|h(t)λεcf0|=|1eλεt+cf0λε(1eλεt)+eλεt0tφ(s)eλεsds1cf0λε|=λεcf0|eλεt(1cf0λε)+eλεt0tφ(s)eλεsds|eλεt+eλεt0t(cf0+φ(s))eλεsdsCˆεeρεt,

where for the last inequality we have used that the denominator is a positive continuous function bounded below (it takes the value 1 for t=0 and its limit is cf0λε when t goes to infinity). We also used the following estimate for the numerator: since, by Lemma 2.7, |φ(s)|Kεeρεsf0, then

|eλεt(1cf0λε)+eλεt0tφ(s)eλεsds|eλεt(|1cf0λε|Kεf0λερε)+Kεf0λερεeρεt2Kεeρεtf0.

 □

In order to give an estimate of the dependence of Cˆε w.r.t. ε, we need to bound the denominator more precisely and to take a value of ρε separated of its limit value. As in Lemma 2.8, we choose ρε=λε(1ε)2=:αε2.

Lemma 2.10

For ε>0 small enough, there exist constants K7 and K8 (independent of ε) such that

|h(t)λεcf0|K8εK7ε2eαεt2.

Proof

Using Lemma 2.7 and the fact that the second term is positive we see that

eλεt+eλεt0t(cf0+φ(s))eλεsdseλεt+max(0,cf0(1eλεt)Kεeρεt)eλεtε (22)

for any tε such that

cf0(1eλεtε)Kεeρεtεeλεtε. (23)

(Notice that the left hand side in (23) is an increasing function of tε.) This indeed happens if Kεeρεtεcf02 and (1+cf0)eλεtεcf02. Since the second condition is weaker than the first one for ε small enough, (23) holds whenever tε is such that eρεtεcf02Kε, i.e., eλεtε(cf02Kε)λερε and ε>0 is sufficiently small. So, (cf02Kε)λερε is also a lower bound in (22), and we finally have

|eλεt+eλεt0t(cf0+φ(s))eλεsds|(cf02Kε)λερε.

Using the bound on the numerator given in the proof of Lemma 2.9, the previous estimate, and using also Lemma 2.8, Lemma 2.6 and Proposition 2.3, we obtain

|h(t)λεcf0|2Kεeρεtf0(cf02Kε)λερε2K5ε4eαεt2f0(K1ε22K5ε4)K6ε2=2K5(2K5K1)K6ε2ε46K6ε2eαεt2f0K8εK7ε2eαεt2. (24)

 □

Finally, a standard application of the triangular inequality and Lemma 2.6, Lemma 2.7, Lemma 2.9 give

f(t,)λεψε(x)|h(t)λεcf0|T˜ε(t)f0+λεcf0T˜ε(t)f0cf0ψε(x)Cˆεeρεt(K2+Kεeρεtf0)+1K1ε2KεeρεtCεeρεt. (25)

Using Lemma 2.8, Lemma 2.10 in the second inequality of (25), the last statement of Proposition 2.4 follows.

2.4. Proof of Theorem 1.2

By the triangular inequality,

f(t,)εγ(0)(γ(0)πε)2+2L1(I)f(t,)λεψεL1(I)+λεψεεγ(0)(γ(0)πε)2+2L1(I).

Hence by Proposition 2.3 and Proposition 2.4, we only need to estimate the last term, for which we have

λεεγλε(1ε)+2εγ(0)(γ(0)πε)2+2L1(I)ε(λε1)γλε(1ε)+2L1(I)+εγλε(1ε)+2εγ(γ(0)πε)2+2L1(I)+ε(γγ(0))(γ(0)πε)2+2L1(I).

Let us bound the three terms. For the first one we have, by Proposition 2.3,

ε(λε1)γλε(1ε)+2L1(I)(λε1)γRεdx(γ(0)πε)22+x2=(λε1)γ2γ(0)=O(ε).

For the second one, by Proposition 2.3 and (17),

εγλε(1ε)+2εγ(γ(0)πε)2+2L1(I)|(γ(0)πε)2(λε(1ε))|εγRdx((γ(0)πε)22+x2)2=|(γ(0)πε)2(λε(1ε))|K0ε2=O(εln1ε).

For the third one, similarly to the proof of Proposition 2.3, denoting by A:=γγ,

ε(γγ(0))(γ(0)πε)2+2L1(I)2ε0Aγx(γ(0)πε)2+x2dx+2εA+γ(γ(0)πε)2+x2dx=εγln(1+A2(γ(0)πε)2)+2γγ(0)πarctan(γ(0)πεA)=O(εln1ε).

2.5. Proof of Lemma 2.8

Let us consider the linear initial value problem

{u(t,x)t=A˜εu(t,x)=(aε(x)λε)u(t,x)+εγ(x)Iu(t,y)dy,u(0,x)=u0(x), (26)

where aε(x):=1εx2. Let us recall that s(A˜ε)=0 and εIγ(x)λεaε(x)dx=1 (see Proposition 2.2). Applying the Laplace transform with respect to t to the previous equation, we obtain the identity

μL[u](μ,x)u0(x)=(aε(x)λε)L[u](μ,x)+εγ(x)IL[u](μ,y)dy,

that is

L[u](μ,x)=u0(x)μ+λεaε(x)+εγ(x)μ+λεaε(x)IL[u](μ,y)dy. (27)

Integrating (with respect to x), we obtain

IL[u](μ,x)dx=Iu0(x)μ+λεaε(x)dx1Iεγ(x)μ+λεaε(x)dx=Iu0(x)μ+λεaε(x)dxεμIγ(x)(λεaε(x))(μ+λεaε(x))dx,

where we have used, for the second equality, εIγ(x)λεaε(x)=1. Substituting in (27), we get

L[u](μ,x)=u0(x)μ+λεaε(x)+Iu0(x)μ+λεaε(x)dxμIγ(x)(λεaε(x))(μ+λεaε(x))dxγ(x)(μ+λεaε(x)). (28)

This Laplace transform is analytic for Re μ>0 (note that λεaε(x) is positive and tends to zero when ε tends to zero). Then, for s>0, we know, by the inversion theorem, that

u(t,x)=12πisis+iL[u](μ,x)eμtdμ.

Using the theorem of residues, we can shift the integration path to the left in order to obtain, for any s(1ελε,0),

u(t,x)=Resμ=0(L[u](μ,x)eμt)+12πisis+iL[u](μ,x)eμtdμ,

where

Resμ=0(L[u](μ,x)eμt)=limμ0μL[u](μ,x)=limμ0(μu0(x)μ+λεaε(x)+Iu0(x)μ+λεaε(x)dxIγ(x)(λεaε(x))(μ+λεaε(x))dxγ(x)μ+λεaε(x))=u0,ψεψε,ψεψε(x)=cu0ψε(x)

(let us recall that ψε(x)=εγ(x)λεaε(x) and ψε(x)=(εIγ(x)dx(λε(1εx2))2)1λε(1εx2)).

Thus, we obtain that, for s(1ελε,0),

u(t,x)=cu0ψε(x)+12π+L[u](s+iτ,x)e(s+iτ)tdτ. (29)

We now define gε(μ):=Iu0(x)dxμ+λεaε(x)μIγ(x)dx(λεaε(x))(μ+λεaε(x)), so that we can write

12π+L[u](s+iτ,x)e(s+iτ)tdτ=12πu0(x)esteiτts+λεaε(x)+iτdτ+12πγ(x)estgε(s+iτ)eiτts+λεaε(x)+iτdτ=e(λεaε(x))tu0(x)+12πγ(x)estgε(s+iτ)eiτts+λεaε(x)+iτdτ, (30)

where we used the estimate s+λεaε(x)>0 and the identity eiτtα+iτdτ=2πeαt (for α>0).

We now would like to find a bound for 12πγ(x)estgε(s+iτ)eiτts+λεaε(x)+iτdτ.

We see that

γ(x)estgε(s+iτ)eiτts+λεaε(x)+iτdτestγsupx|gε(s+iτ)eiτts+λεaε(x)+iτdτ|=estsupx|gε(s+iτ)eiτts+λεaε(x)+iτdτ| (31)

and

|gε(s+iτ)eiτts+λεaε(x)+iτdτ||gε(s+iτ)||s+λεaε(x)+iτ|dτ|gε(s+iτ)||s+λε(1ε)+iτ|dτ (32)

since |s+λεaε(x)+iτ||s+λε(1ε)+iτ|.

Let us then find an upper bound for gε(s+iτ). For the numerator of gε(s+iτ) we can estimate

|Iu0(x)s+iτ+λεaε(x)dx|u01|s+iτ+λε(1ε)|.

We now find a lower bound for the denominator of gε(s+iτ). We use the elementary estimate |z|max(|Rez|,|Imz|) and we start with the real part.

|ReIγ(x)(λεaε(x))(s+iτ+λεaε(x))dx|=|Iγ(x)(λεaε(x))s+λεaε(x)|s+iτ+λεaε(x)|2dx|=Iγ(x)(λεaε(x))s+λεaε(x)|s+iτ+λεaε(x)|2dx=Iγ(x)(λεaε(x))(s+λεaε(x)+τ2s+λεaε(x))dxIγ(x)(λε0(1ε0)+x2)(λε0(1ε0)+x2+τ2x2)dx=Ix2γ(x)(λε0(1ε0)+x2)((λε0(1ε0)+x2)x2+τ2)dx=:F(τ),

where in the last inequality we used the estimates s+λεaε(x)<λεaε(x), s+λε(1ε)>0. We also used that, since λε(1ε) is strictly positive and tends to zero when ε goes to zero, there exists ε0 such that ε<ε0 we have λε0(1ε0)>λε(1ε).

In a similar way, for the imaginary part,

|ImIγ(x)(λεaε(x))(s+iτ+λεaε(x))dx|=|Iγ(x)(λεaε(x))τ(s+λεaε(x))2+τ2dx|=|τ|Iγ(x)(λεaε(x))((s+λεaε(x))2+τ2)dx|τ|Iγ(x)(λε0(1ε0)+x2)((λε0(1ε0)+x2)2+τ2)dx=:G(τ).

Defining H(τ):=max(F(τ),G(τ)) we see that

|gε(s+iτ)|u01|s+iτ+λε(1ε)||s+iτ|H(τ), (33)

and then, using (31), (32) and (33)

||γ(x)est+gε(s+iτ)eiτts+λεaε(x)+iτdτ||est+dτs2+τ2|s+iτ+λε(1ε)|2H(τ)u01.

Now, since F and G are strictly positive continuous functions, F(0)>0 and τG(τ) tends to a positive limit when τ goes to ∞, there exists a constant C>0 (independent of ε) such that H(τ)C1+τ. Choosing s=αε2, where αε=λε(1ε), we can write

||xγ(x)est+gε(s+iτ)eiτts+λεaε(x)+iτdτ||eαεtC0+2(1+τ)((αε2)2+τ2)32dτu01=eαεt2C(8αε2+4αε)u01.

Finally, going back to (29) and using (30), we end up with

u(t,)cu0ψε(1+1πC(4αε2+2αε))eαεt2u01K5ε4eαεt2u01.

3. Proof of Theorem 1.3

We start here the proof of Theorem 1.3. From now on, C will designate a strictly positive constant depending only on some upper bounds on γL, f0W1,, on a lower bound on f0, and on |ba| (assumed to be finite in this theorem).

Thanks to the variation of the constant formula, the solution f of (1) satisfies:

f(t,x)=f(0,x)e(1εx2)t0tIf(s,y)dyds+ε0t(γ(x)If(s,y)dy)e(1εx2)(ts)stIf(σ,y)dydσds=f(0,x)e(1εx2)t0tI(s)ds+ε0tγ(x)I(s)e(1εx2)(ts)stI(σ)dσds, (34)

where

I(t):=If(t,y)dy.

Obtaining a precise estimate on te(1ε)(ts)stI(σ)dσ is the cornerstone of the proof of Theorem 1.3.

3.1. Preliminary estimates

If we sum (34) along xR, we get, for t0:

I(t)=(If(0,x)ex2tdx)e(1ε)t0tI(s)ds+ε0t(IIγ(x)f(s,y)ex2(ts)dxdy)e(1ε)(ts)stI(σ)dσds=z1(t)te(1ε)t0tI(s)ds+ε0tz2(s,ts)tse(1ε)(ts)stI(σ)dσds, (35)

where

z1(t):=tIf(0,x)ex2tdx,z2(σ,τ)=τIIγ(x)f(σ,y)ex2τdxdy.

If we integrate our equation w.r.t. x, we get

It(t)=I(t)(1εI(t))Ix2f(t,x)dx+εIIγ(x)f(t,y)dxdyI(t)(1εI(t))+εI(t)I(t)(1I(t)),

which implies, since I(0)1, that

0I(t)1. (36)

Thanks to (35), (36) and the nonnegativity of z1,z2, one gets

z1(t)te(1ε)t0tI(s)dsC, (37)

while for some constants C,C>0,

z1(t)=If(0,xt)ex2dx1CCCf(0,xt)dx1C,

for t1. Note that here we used a lower bound on f(0,) around x=0 (we have assumed that f(0,0)>0 and that f(0,) is continuous). Thanks to this lower bound, (37) becomes

e(1ε)t0tI(s)dsCt. (38)

Thanks to (38) and (36), we can estimate the second term of (35) as follows:

w(t):=ε0tz2(s,ts)tse(1ε)t0tI(σ)dσeεs+0s(I(σ)1)dσdsCεtz2L0teCεstsdsCεtz2LeCεt0teCεssdsCεtz2LeCεt. (39)

In order to estimate z2L, we proceed as follows:

z2(s,τ)=IIγ(xτ)f(s,y)ex2dxdyCI(s)Iex2dxC.

This estimate combined with (39) implies that w(t) satisfies

0w(t)CεteCεt. (40)

Since f(0,)W1,(I), we can estimate (for t>0)

z1(t)=I(f(0,0)+0xtfx(0,z)dz)ex2dx=f(0,0)Iex2dx+λ(t), (41)

where

|λ(t)|I|0xtfx(0,z)dz|ex2dxCtI|x|ex2dxCt. (42)

3.2. Estimate for e(1ε)t0tI(s)ds

Thanks to (35) (and the definition of λ and w: see (41) and (39) respectively), we see that, for t1,

I(t)=f(0,0)Iex2dx+λ(t)te(1ε)t0tI(s)ds+w(t), (43)

so that

e0tI(s)ds=e01I(s)ds+1tdds(e0sI(σ)dσ)(s)ds=e01I(s)ds+1tf(0,0)Iex2dxse(1ε)sds+1tλ(s)se(1ε)sds+1tw(s)e0sI(σ)dσds. (44)

We will now estimate each of the terms on the right hand side of (44). We start by estimating the third term on the right hand side, thanks to (42) and an integration by parts:

|1tλ(s)se(1ε)sds|C1te(1ε)ssdsC[e(1ε)t(1ε)t+1te(1ε)s(1ε)s2ds]C1ε[e(1ε)tt+tmaxs[1,t]e(1ε)ss2]2C(1ε)te(1ε)t, (45)

provided that t>4 (this last assumption ensures, remembering that 0<ε<1/2, that maxs[1,t]e(1ε)ss2=e(1ε)tt2).

We now estimate the second term on the right hand side of (44), using an integration by parts:

1tf(0,0)Iex2dxse(1ε)sds=f(0,0)(Iex2dx)(e(1ε)t(1ε)te1ε1ε+1te(1ε)s2(1ε)s3/2ds),

and then, applying an estimate similar to the one used to obtain (45), we get, provided that t>4,

01te(1ε)s2(1ε)s3/2ds1te(1ε)s2(1ε)sds1(1ε)2te(1ε)t. (46)

Finally, we estimate the last term of the right hand side of (44), thanks to estimates (40) and (36), and for t1:

01tw(s)e0sI(σ)dσds1t|w(s)|eIL(R+)sdsCε1tseCεsesdsCεte(1+Cε)t1+Cε, (47)

where we have used st to obtain the last inequality.

Combining these estimates, estimate (44) becomes:

e0tI(s)ds(1ε)t=f(0,0)Iex2dx(1ε)t+μ(t), (48)

or

e(1ε)t0tI(s)ds=(f(0,0)Iex2dx(1ε)t+μ(t))1, (49)

where, thanks to (44), (45), (46) and (47), for t>4,

Ctμ(t)C(1t+εteCεt). (50)

3.3. Estimate for |e(1ε)t0tI(s)dstf(0,0)Iex2dx|

Thanks to (49),

|e(1ε)t0tI(s)dstf(0,0)Iex2dx|=|(f(0,0)Iex2dx(1ε)t+μ(t))1tf(0,0)Iex2dx| (51)
=tf(0,0)Iex2dx|(11ε+μ(t)tf(0,0)Iex2dx)11|. (52)

We notice that thanks to estimate (50),

f(0,0)Iex2dx+(1ε)μ(t)tf(0,0)Iex2dx2, (53)

as soon as t>4.

Under the same assumption, we directly get from (50) that

|μ(t)|C(1t+εteCεt). (54)

Using the bounds (53) and (54), we can show that as soon as t>4,

|(11ε+μ(t)tf(0,0)Iex2dx)11|=|εf(0,0)Iex2dx(1ε)μ(t)tf(0,0)Iex2dx+(1ε)μ(t)t|C(1t+εt32eCεt), (55)

so that identity (52) leads to the bound (for t>4)

|e(1ε)t0tI(s)dstf(0,0)Iex2dx|C(1+εt2eCεt). (56)

Notice also, as this is going to be useful further on, that for s>4, thanks to (48) and (54),

|e0sI(σ)dσ(1ε)sf(0,0)Iex2dxs|=|μ(s)+εf(0,0)Iex2dx(1ε)s|C(1s+εseCεs). (57)

3.4. Conclusion of the proof of Theorem 1.3

In this last part of the proof, we systematically consider times t>4. We estimate

xf(t,x)f(0,x)tex2tf(0,0)Iey2dyL1(I)xf(t,x)f(0,x)ex2tf(0,0)Iey2dy(1ε)t+μ(t)L1(I)+xf(0,x)ex2tf(0,0)Iey2dy(1ε)t+μ(t)f(0,x)tex2tf(0,0)Iey2dyL1(I). (58)

Let us start by estimating the second term of the right hand side of (58), thanks to estimate (55):

xf(0,x)ex2tf(0,0)Iey2dy(1ε)t+μ(t)f(0,x)tex2tf(0,0)Iey2dyL1(I)xf(0,x)tex2tf(0,0)Iey2dy|(11ε+μ(t)tf(0,0)Iey2dy)11|L1(I)f(0,)tf(0,0)Iey2dy|(11ε+μ(t)tf(0,0)Iey2dy)11|Iex2tdxC(1t+εt32eCεt). (59)

We now rewrite the first term of the right hand side of (58), using formula (34) and (49):

xf(t,x)f(0,x)ex2tf(0,0)Iey2dy(1ε)t+μ(t)L1(I)=xε0t(Iγ(x)f(s,y)dy)e(1εx2)(ts)stI(σ)dσdsL1(I)CεI0tI(s)ex2(ts)e(1ε)t0tI(σ)dσe0sI(σ)dσ(1ε)sdsdx,

and then, thanks to (36), (56) and (57),

xf(t,x)f(0,x)ex2tf(0,0)Iey2dy(1ε)t+μ(t)L1(I)Cε04(Iex2(ts)dx)(tf(0,0)Iex2dx+1+εt2eCεt)ds+Cε4t(Iex2(ts)dx)(f(0,0)Iex2dxs+1s+εseCεs)dqsfesrgqdreg(tf(0,0)Iex2dx+1+εt2eCεt)dsCε1t(t+1+εt2eCεt)+Cε4t1ts(1s+εseCεs)(t+1+εt2eCεt)ds.

We estimate

4tseCεstsdsteCεt4tdstsCt32eCεt,

and then

xf(t,x)f(0,x)ex2tf(0,0)Iey2dy(1ε)t+μ(t)L1(I)Cε(1+1t+εt32eCεt)+Cε(1+εt32eCεt)(t+1+εt2eCεt)C(ε+εt+εt+(εt32+ε2t2+ε2t32+ε3t72)eCεt)C(εt+εt32eCεt), (60)

where we have used the fact that εtCeCεt. Thanks to (59) and (60), (58) becomes:

xf(t,x)f(0,x)tex2tf(0,0)Iey2dyL1(I)C(1t+εt+εt32eCεt).(1t+εt32)eCεt.

Theorem 1.3 follows from this estimate.

Note that as stated in the comments of the theorem, if we assume that t[1κ2,κ23ε23], then (6) becomes

xf(t,x)f(0,x)tex2tf(0,0)Iey2dyL1(I)C(κ+κeCκ23ε13),

and if furthermore εκ1, then

xf(t,x)f(0,x)tex2tf(0,0)Iey2dyL1(I)Cκ.

Acknowledgments

The research leading to this paper was funded by the French “ANR blanche” project Kibord: ANR-13-BS01-0004, and “ANR JCJC” project MODEVOL ANR-13-JS01-0009, and by the Idex program of Université Sorbonne Paris Cité, in the framework of the “Investissements d'Avenir”, convention ANR-11-IDEX-0005. A.C. and S.C. were partially supported by the Ministerio de Economía y Competitividad, Grants MTM2011-27739-C04-02 and MTM2014-52402-C3-2-P.

Submitted by Y. Du

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